Richard Connor


Richard Connor


University of Michigan
Evolutionary Biology

Phone: 508-999-8221 
Office: Violette Research Building, Room 215

For the past 30 years I have worked on a unique population of Indian Ocean bottlenose dolphins (Tursiops aduncus) in Shark Bay, Western Australia. Most of my work has focused on alliances formed by adult males in competition over estrus females. These alliances are remarkable for their complexity. The males form three levels of nested alliances, a phenomenon that has no match outside of our own species. Our findings have important implications for current thinking about the role complex social relationships had in the evolution of large brain size and intelligence in mammals. We are working on over 100 adult males in the population and are launching a fully integrated study that examines alliance formation from multiple perspectives, including communication, cognition, genetics, ecology, and development.

In addition to working in Shark Bay, my graduate students have worked on dolphins and whales in a variety of locations in the US, Central America, the Caribbean and India, exploring a range of questions focused on behavior and ecology.

I also have a long-standing interest in theory, especially on the evolution of cooperation, altruism and mutualism and the evolution of social systems.

 Recent Publications | Courses



For a complete list of Dr. Connor's publications, see Google Scholar 

For more project information and updates on the Dolphin Alliance Project visit our Facebook page


Dolphins & Whales

Connor, R.C. &Krützen, M. (2015) Male dolphin alliances in Shark Bay: changing perspectives in a thirty-year study. Animal Behavior, 103: 223-235.

Connor, R.C., J.J. Watson-Capps, W.B. Sherwin & M. Krützen. 2011 New levels of complexity in the male alliance networks of Indian Ocean bottlenose dolphins (Tursiops sp.). Biology Letters 7,623-626

Connor, R.C. 2007. Complex alliance relationships in bottlenose dolphins and a consideration of selective environments for extreme brain size evolution in mammals. Philosophical Transactions of the Royal Society: Biological Sciences 362: 587-602.

Morisaka, T. & Connor, R.C. 2007. Predation by killer whales (Orcinus orca) and the evolution of whistle loss and narrow-band high frequency clicks in odontocetes. J. Evol. Biol. 20: 1439-1458

Connor, R.C., Smolker, R.A. & L.Bejder. 2006. Synchrony, social behavior and alliance affiliations in Indian Ocean bottlenose dolphins (Tursiops aduncus). Animal Behavior. 72:1371-1378.

Connor, R.C., Mann, J., & J.-Capps. 2006. A sex-specific affiliative contact behavior in Indian Ocean bottlenose dolphins, Tursiops sp. Ethology 112: 631-638.

Krützen, M., J. Mann, M. R. Heithaus, R. C. Connor, L. Bejder & W. B.Sherwin. 2005. Cultural transmission of a foraging strategy involving tool use in bottlenose dolphins. Proceedings of the National Academy of Sciences. 102(25): 8939-8943.

Connor, R.C., Heithaus, M.R. & Barre, L.M. 2001. Complex social structure, alliance stability and mating access in a bottlenose dolphin 'super-alliance'. Proceedings of the Royal Society of London, Biological Sciences. 268: 263-267.

Connor, R.C. 2001. Individual foraging specializations in marine mammals: culture and ecology. Behavioral and Brain Sciences, 24:329-330

Connor, R.C., R. Wells, J. Mann, and A. Read. 2000. The bottlenose dolphin: social relationships in a fission-fusion society. In: Cetacean Societies: Field studies of whales and dolphins, eds: J. Mann; R.Connor, P.Tyack, and H. Whitehead. University of Chicago Press, pp 91-126.

Connor, R.C.; Heithaus, R.M. and Barre, L.M. 1999. Super-alliance of bottlenose dolphins. Nature, 371: 571-572

Corkeron, P.J. and R.C. Connor. 1999. Why do baleen whales migrate? Marine Mammal Science. 15: 1228-1245.

Connor, R.C.; Mann, J; Tyack, P.L. and Whitehead, H. 1998. Social evolution in toothed whales. Trends in Ecology & Evolution 13: 228-232

Connor, R.C., Richards, A.F., Smolker, R.A. & J.Mann. 1996. Patterns of female attractiveness in Indian Ocean bottlenose dolphins. Behaviour 133: 37-69.

Connor, R.C., Smolker, R.A., & A.F. Richards. 1992. Two levels of alliance formation among male bottlenose dolphins (Tursiops sp.). Proc. Nat. Acad. Sci.. 89: 987-990.


The evolution of cooperation & mutualism

Connor, R.C. 2010. Cooperation beyond the dyad: from simple models to a complex society. Philosophical Transactions of the Royal Society: Biological Sciences. 365:2687-2697

Connor, R.C. 2007. Invested, extracted & byproduct benefits: a modified scheme for the evolution of cooperation. Behav. Proc. 76: 109-113.

Whitehead, H. & R.C. Connor 2005. Alliances I: How large should alliances be? Animal Behaviour 69: 117-126.

Connor, R.C. & H. Whitehead. 2005.Alliances II: Rates of encounter during resource utilization: A general model of intrasexual alliance formation in fission-fusion societies. Animal Behaviour 69: 127-132.

Liemar, O. & R. Connor. 2003. Byproduct benefits, reciprocity and pseudo-reciprocity in mutualism. In: Hammerstein, P., ed. Genetic & Cultural Evolution of Cooperation, Dahlem Workshop Report 90, MIT Press; pp. 203-22

Connor, R.C. 1996. Partner preferences in by-product mutualisms and the case of predator inspection in fish. Animal Behaviour 51: 451-454.

Connor, R.C. 1995. Altruism among non-relatives: alternatives to the Prisoner's Dilemma. Trends in Ecology & Evolution, 10: 84-86.

Connor, R.C. & Curry R. 1995. Helping non-relatives: a role for deceit? Anim. Behav., 49: 389-393.

Connor, R.C. 1995. The benefits of mutualism: a conceptual framework. Biol. Rev., 70: 427-457.

Connor, R.C. 1995. Impalla allogrooming: tit-for-tat or parcelling? Anim. Behav., 49: 528-530.

Connor, R.C. 1992 Egg-trading in simultaneous hermaphrodites: An alternative to tit-for-tat. J. Evol. Biol . 5: 523-528.

Connor, R.C. 1986. Pseudoreciprocity: Investing in mutualism. Animal Behaviour 34: 1562-1566.


Graduate Student Publications

Gisburne, T.J. & Connor, R.C. 2015. Group size and feeding success in strand-feeding bottlenose dolphins (Tursiops truncatus) in Bull Creek, South Carolina. Marine Mammal Science. DOI: 10.1111/mms.12207

Gazda, S.Swami Iyer,S.; Killingback, T.; Connor, R., & Brault, S. 2015.The Importance of Delineating Networks by Activity Type in Bottlenose Dolphins (Tursiops truncatus) in Cedar Key, Florida. Royal Society Open Science

Eierman, L.E. &  Connor R.C. 2014. Foraging behavior, prey distribution and microhabitat use by bottlenose dolphins (Tursiops truncatus) in a tropical atoll.  Marine Ecology Progress Series, 503:279-288. doi:10.3354/meps10721

Randic, S, Connor, RC, Sherwin, WS & Krutzen, M. 2012, A novel mammaliansocial in Indo-Pacific bottlenose dolphins (Tursiops sp.): complex male alliances in an open social network. Proc. Roy. Soc. Biol. Sci. 279: 3083-3090.

Connor, R.C. & Vollmer, N. 2009. The puzzle of sexual coercion in chimpanzees. Sexual Coercion in Primates: An Evolutionary Perspective on Male Aggression Against Females( Eds. M. N. Muller and R.W. Wrangham).Cambridge: Harvard University Press.

Duffy-Echevarria, E., Connor, R.C. & D. St. Aubin. 2008. Observations of strand feeding by bottlenose dolphins (Tursiops truncatus) in Bull Creek, South Carolina. Marine Mammal Science, 24:202-206.

Kaplan, J.D. & Connor, R.C. 2007. A preliminary examination of sex differences in tactile interactions among juvenile Atlantic spotted dolphins (Stenella frontalis). Marine Mammal Science, 23: 943-953.

Gazda, S., R.C. Connor, R.Edgar, & F. Cox. 2005. A Division of Labor with Role Specialization in Group-Hunting Bottlenose Dolphins (Tursiops truncatus) off Cedar Key, Florida The Proceedings of the Royal Society:Biological Sciences 272: 135-140.

Sardi, K., Weinrich, M., & R.C. Connor. 2005.Social interactions of humpback whale (Megaptera novaeangliae): Mother calf pairs on a north Atlantic feeding ground. Behaviour.142(6): 731-750.

Sutaria, D., and Jefferson, T. A. (2004). Records of Indo-Pacific Humpback Dolphin (Sousa chinensis Osbeck, 1765) along the coast of India and Sri Lanka - An Overview. Aquatic Mammals Special Issue 30(1) 125-136

Fedorowicz, S.M., Beard, D. & R.C. Connor. 2003. Food sharing in wild bottlenose dolphins. Aquatic Mammals. 29.3: 355-359.

Connor, R.C., Heithaus, M.R., Berggren, P. and J.L. Miksis. 'Kerplunking': Surface fluke-splashes during shallow water bottom foraging by bottlenose dolphins. 2000. Mar. Mamm. Sci. 16: 646-653.

Miksis, J.L., Grund, M.D., Nowacek, D.P., Connor, R.C. & Tyack, P.L. 2001. Cardiac Responses to Acoustic Playback Experiments in the Captive Bottlenose Dolphin, Tursiops truncatus. J. Comp. Psych. 115: 227-332.


Courses Professor Connor has taught include:

  • Topics in Biology – Marine Mammals (BIO 103)
  • Biology of Populations (BIO 210)
  • Biology of Marine Mammals (BIO 414)
  • Animal Behavior (BIO 420)
  • Cooperation and Mutualism (BIO411)
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